About the Authors

Wei Sun

Contributed equally to this work with: Wei Sun, Katerina Kechris, Sean Jacobson

Current address: Program in Biostatistics and Biomathematics, Public Health Sciences Division, Fred Hutchinson Cancer Research Center, Seattle, Washington, United States of America

Affiliation Department of Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

Katerina Kechris

Contributed equally to this work with: Wei Sun, Katerina Kechris, Sean Jacobson

Affiliation Department of Biostatistics and Informatics, Colorado School of Public Health, University of Colorado Anschutz Medical Campus, Aurora, Colorado, United States of America

Sean Jacobson

Contributed equally to this work with: Wei Sun, Katerina Kechris, Sean Jacobson

Affiliation National Jewish Health, Denver, Colorado, United States of America

M. Bradley Drummond

Affiliation Division of Pulmonary, Critical Care and Sleep Medicine, Department of Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland, United States of America

Gregory A. Hawkins

Affiliation Center for Genomics and Personalized Medicine Research, Wake Forest School of Medicine, Winston-Salem, North Carolina, United States of America

Jenny Yang

Affiliation Department of Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

Ting-huei Chen

Affiliation Department of Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

Pedro Miguel Quibrera

Affiliation Collaborative Studies Coordinating Center, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

Wayne Anderson

Affiliation Marsico Lung Institute/Cystic Fibrosis Research Center, Department of Medicine, Division of Pulmonary and Critical Care Medicine, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina United States of America

R. Graham Barr

Affiliation Department of Medicine, Columbia University Medical Center, New York, New York; Department of Epidemiology, Mailman School of Public Health at Columbia University, New York, New York, United States of America

Patricia V. Basta

Affiliation Department of Epidemiology, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

Eugene R. Bleecker

Affiliation Center for Genomics and Personalized Medicine Research, Wake Forest School of Medicine, Winston-Salem, North Carolina, United States of America

Terri Beaty

Affiliation Department of Epidemiology, Bloomberg School of Public Health, Johns Hopkins University,Baltimore, Maryland, United States of America

Richard Casaburi

Affiliation Division of Respiratory and Critical Care Physiology and Medicine, Harbor- University of California at Los Angeles Medical Center, Torrance, California, United States of America

Peter Castaldi

Affiliation Channing Division of Network Medicine, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America

Michael H. Cho

Affiliation Channing Division of Network Medicine, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America

Alejandro Comellas

Affiliation Division of Pulmonary and Critical Care Medicine, University of Iowa, Iowa City, Iowa, United States of America

James D. Crapo

Affiliation Department of Medicine, Division of Pulmonary, Critical Care and Sleep Medicine, National Jewish Health, Denver, Colorado, United States of America

Gerard Criner

Affiliation Department of Thoracic Medicine and Surgery, Lewis Katz School of Medicine, Temple University, Philadelphia, Pennsylvania, United States of America

Dawn Demeo

Affiliation Division of Pulmonary and Critical Care Medicine, Channing Division of Network Medicine, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America

Stephanie A. Christenson

Affiliation Division of Pulmonary, Critical Care, Allergy, and Sleep Medicine, Department of Medicine, University of San Francisco Medical Center, University of California San Francisco, San Francisco, California, United States of America

David J. Couper

Affiliation Collaborative Studies Coordinating Center, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

Jeffrey L. Curtis

Affiliation Division of Pulmonary and Critical Care Medicine, University of Michigan Health System, Ann Arbor, Michigan; VA Ann Arbor Healthcare System, Ann Arbor, Michigan, United States of America

Claire M. Doerschuk

Affiliation Marsico Lung Institute/Cystic Fibrosis Research Center, Department of Medicine, Division of Pulmonary and Critical Care Medicine, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina United States of America

Christine M. Freeman

Affiliation Division of Pulmonary and Critical Care Medicine, University of Michigan Health System, Ann Arbor, Michigan; VA Ann Arbor Healthcare System, Ann Arbor, Michigan, United States of America

Natalia A. Gouskova

Current address: Harvard T.H. Chan School of Public Health, Boston, Massachusetts, United States of America

Affiliation Collaborative Studies Coordinating Center, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

MeiLan K. Han

Affiliation Division of Pulmonary and Critical Care Medicine, University of Michigan Health System, Ann Arbor, Michigan, United States of America

Nicola A. Hanania

Affiliation Section of Pulmonary and Critical Care Medicine, Baylor College of Medicine, Houston, Texas, United States of America

Nadia N. Hansel

Affiliation Division of Pulmonary and Critical Care Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland, United States of America

Craig P. Hersh

Affiliation Channing Division of Network Medicine, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America

Eric A. Hoffman

Affiliation Department of Radiology, Division of Physiologic Imaging, University of Iowa Hospitals and Clinics, Iowa City, Iowa, United States of America

Robert J. Kaner

Affiliation Department of Genetic Medicine, Weill Cornell Medical College, New York, New York, Department of Medicine, Division of Pulmonary and Critical Care Medicine, Weill Cornell Medical College, New York, New York, United States of America

Richard E. Kanner

Affiliation Department of Internal Medicine, Division of Pulmonary and Critical Care Medicine, University of Utah, Salt Lake City, Utah, United States of America

Eric C. Kleerup

Affiliation Division of Pulmonary and Critical Care Medicine, Department of Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Sharon Lutz

Affiliation Department of Biostatistics and Informatics, Colorado School of Public Health, University of Colorado Anschutz Medical Campus, Aurora, Colorado, United States of America

Fernando J. Martinez

Affiliation Department of Medicine, Weill Cornell Medical College, New York-Presbyterian Hospital/Weill Cornell Medical Center, New York, New York, United States of America

Deborah A. Meyers

Affiliation Center for Genomics and Personalized Medicine Research, Wake Forest School of Medicine, Winston-Salem, North Carolina, United States of America

Stephen P. Peters

Affiliation Department of Medicine, Division of Pulmonary, Critical Care, Allergy and Immunologic Medicine, Wake Forest University School of Medicine, Winston-Salem, North Carolina, United States of America

Elizabeth A. Regan

Affiliation Department of Medicine, National Jewish Health, Denver, Colorado United States of America

Stephen I. Rennard

Current address: Early Clinical Development, Innovative Medicines, AstraZeneca, Melbourn, Hertfordshire, England

Affiliation Division of Pulmonary and Critical Care Medicine, University of Nebraska, Omaha, Nebraska, United States of America

Mary Beth Scholand

Affiliation Department of Internal Medicine, Division of Pulmonary and Critical Care Medicine, University of Utah, Salt Lake City, Utah, United States of America

Edwin K. Silverman

Affiliation Channing Division of Network Medicine, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America

Prescott G. Woodruff

Affiliation Division of Pulmonary, Critical Care, Sleep and Allergy, Department of Medicine and Cardiovascular Research Institute, University of California San Francisco School of Medicine, San Francisco, California, United States of America

Wanda K. O’Neal

wanda_o’neal@med.unc.edu (WKO); RBowler@NJHealth.edu (RPB)

Affiliation Marsico Lung Institute/Cystic Fibrosis Research Center, Department of Medicine, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina United States of America

Russell P. Bowler

wanda_o’neal@med.unc.edu (WKO); RBowler@NJHealth.edu (RPB)

Affiliation Department of Medicine, Division of Pulmonary Medicine, National Jewish Health, Denver, Colorado, United States of America

SPIROMICS Research Group

Membership in SPIROMICS and COPDGene is provided in the Acknowledgments.

COPDGene Investigators

Membership in SPIROMICS and COPDGene is provided in the Acknowledgments.

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: RPB WKO KK WS. Performed the experiments: RPB MBD GAH WKO. Analyzed the data: RPB ThC SJ KK WKO WS JY. Contributed reagents/materials/analysis tools: WA RGB PVB ERB TB RC PC MHC AC JDC GC DD SAC DJC JLC CMD MBD CMF NAG MKH NAH NNH GAH CPH EAH RJK REK ECK SL FJM DAM SPP PMQ EAR SIR MBS EKS PGW. Wrote the paper: RPB KK WKO WS.